Avoidance of parasitoid attack is associated with the spatial use within a leaf by a lepidopteran leafminer

In prey‐predator systems, top‐down effects can be a powerful determinant for spatial distributions of prey through their search for enemy‐free space. Leafminers live and eat within leaves, making feeding tracks called mines, and mine conspicuousness exposes them to a high risk of parasitism. Those lepidopteran leafminers that use lower leaf surfaces as mining sites show wide evolutionary radiation. We hypothesized that leafminers making mines on the lower surface are less often detected by parasitoids and thus have a selective advantage in avoiding parasitism compared to those on the upper surface. To investigate the adaptiveness of lower‐surface mining, we examined the relationship between parasitism and within‐leaf mine distribution for 3 years using a field population of the leafminer Phyllocnistis spec. (Lepidoptera: Gracillariidae, Phyllocnistinae), which prefers the lower surface of leaves of the Japanese privet, Ligustrum japonicum Thunb. (Oleaceae). Parasitoid attack was more frequent in the upper‐surface mines than in the lower‐surface mines and on leaves with upper‐surface mines than on leaves with only lower‐surface mines. When both surfaces were mined, leafminers on the lower surface could avoid parasitism. Upper‐surface mines were attacked by more parasitoid species as compared to lower‐surface mines. Although the results demonstrated that mining on the lower surface was advantageous in avoiding parasitism, the vulnerability of lower‐surface mines to parasitism varied depending on their abundance. When many lower‐surface mines were present, lower‐surface mines suffered a higher parasitism rate than upper‐surface mines, probably because parasitoids formed search images for and concentrated on lower‐surface mines. This study suggests that the preferential use of the lower leaf surface by leafminers is in part attributed to interactions with parasitoids.     

Does the Existence of Bird's Nest Ferns Enhance the Diversity of Oribatid (Acari: Oribatida) Communities in a Subtropical Forest?

We examined the effects of the presence of bird's nest ferns on the species diversity of oribatid mites in the whole forest in terms of the three categories of species diversity (α-, β-, and γ-diversity) in a subtropical forest in south-western Japan. The species diversity (1 − D) of oribatid communities in the ferns was significantly lower than those in bark of trees and the forest-floor litter and soil, and was similar to that in the branches. The oribatid faunas in the litter in and the roots of the fern were more similar to those in both the forest-floor litter and soil than to the faunas in the other arboreal habitats. However, the ferns can be colonized by endemic oribatid species specialized to such environments. The number of oribatid species estimated for a hypothetical stand with no ferns was about 180 species from 80 samples; this value did not differ significantly from that in another hypothetical stand with ferns (ca. 190 species). Thus, the species richness of oribatid communities estimated for the whole forest (the γ-diversity) was not affected by the presence or absence of bird's nest ferns. The α- and β-diversities of oribatid communities on bird's nest ferns were lower than those in other habitats, and they might not dramatically raise the overall γ-diversity of invertebrate communities in the whole forest. The bird's nest ferns, however, can generate a unique habitat for specialized species, and this would help to maintain species diversities of invertebrates at the whole-forest scale in subtropical forests.     

Spatiotemporal distribution of fruitbodies of ectomycorrhizal fungi in an Abies firma forest

 Spatial associations between ectomycorrhizal (ECM) fungi and their presumed host trees, and spatiotemporal associations among ECM fungi were surveyed for 3 years in an Abies firma-dominated forest in central Japan. A total of 39 species in 13 genera of ECM fungi were recorded, with more species in the Russulaceae than any other family. Russula ochroleuca, Russula sp.1 and Strobilomyces confusus tended to produce their fruitbodies on the forest floor directly under the crown of A. firma, whereas those of Inocybe cincinnata, Gomphus floccosus and G. fujisanensis were aggregated in limited areas outside the A. firma crown. Interspecific spatial associations were analysed for Russula sp.1, which was the most dominant species, and three other frequent species, I. cincinnata, S. confusus and R. ochroleuca. Pairwise, Russula sp.1 with I. cincinnata, with S. confusus or with R. ochroleuca showed an association which was exclusive, overlapping or independent, respectively. Fruiting phenologies differed in that S. confusus showed a peak density in the summer, whereas the other three species peaked in the autumn. These results suggest that the formation of ECM fruitbodies can be partitioned among the species both spatially and temporally. Accepted: 7 July 1998     

Resource utilization and sex allocation in response to host size in two ectoparasitoid wasps on subcortical beetles

Patterns of host resource utilization and sex ratio manipulation in relation to host size were investigated for two solitary ectoparasitoid wasps,Atanycolus initiator andSpathius brevicaudis (Hymenoptera Braconidae). Both species parasitize subcortical beetles on the trunks of Japanese pine trees.A. initiator is on average 8 times larger in body weight and has an ovipositor that is 3.7 times longer than that ofS. brevicaudis. In both parasitoids, the size of emerging wasps was positively correlated with host size, but the host/wasp size regressions were linear for all three major host species inA. initiator, whereas inS. brevicaudis the regression was logarithmic for a relatively large host species. The sex ratios (proportion of males) of both parasitoids emerging from different host species decreased with increasing host size, but the overall sex ratio on each host species was male-biased inA. initiator, while female-biased inS. brevicaudis. How the proportion of host consumed changed in response to host size, differed between the two parasitoids for the same host species. In the field survey, the size and sex ratio of the emerging two parasitoids from a dead tree were closely related to host size. However, the spatial distribution of the two parasitoids depended on the bark thickness of the trunk. The data suggest that differences in the relative evaluation of host size and in ovipositor length may enable the coexistence of the two parasitoid wasps.     

Reproductive strategy of a woodwasp with no fungal symbionts, Xeris spectrum (Hymenoptera: Siricidae)

Experiments were conducted to elucidate the reproductive strategy of the siricid woodwasp, Xeris spectrum, which carries no substantial symbiotic fungi in its body, in a comparison with the life cycles of two fungus-carrying siricid woodwasp species, Sirex nitobei and Urocerus japonicus, by considering ecological traits such as seasonal patterns of occurrence, spatial distribution of emergence on a tree, and oviposition activities. Part of the X. spectrum populations emerged in spring, during May and June, while others emerged in summer, during August and September, simultaneously with other siricid fungus-carrying woodwasps. The vertical distribution pattern of X. spectrum emergence holes on the trunk closely coincided with the emergence hole pattern of S. nitobei. X. spectrum laid few eggs on fresh logs, old logs, or on logs inoculated with potato dextrose agar, whereas on logs inoculated with Amylostereum chailletii or A. areolatum, X. spectrum females oviposited no less than 30%, on average, of their potential eggs. Moreover, the oviposition sites on these logs were concentrated near the Amylostereum inoculation positions. These results indicate that X. spectrum has evolved a life history that utilizes fungal symbionts of other woodwasp species without itself possessing any symbiotic fungus. Moreover, X. spectrum has evolved a dual reproductive system in that (1) some adults emerge in summer during the same emergence period as the fungus-carrying woodwasps and thereby oviposit on host trees already inoculated with fungi, and (2) other adults emerge the next spring and oviposit on trees that were inoculated with fungi a year earlier. Received: 16 January 1997 / Accepted: 14 July 1997     

Arthropod communities in a Japanese cedar (Cryptomeria japonica D. Don) plantation: Abundance,biomass and some properties

Arboreal arthropod communities were censused by insecticidal knockdown in a plantation of Japanese cedar,Cryptomeria japonica, in central Japan from April 1983 to February 1985 at intervals of two months. All arthropods sampled were allocated to higher taxonomic groups or guilds and the composition of the communities was analyzed with respect to abundance, biomass and their seasonal trends. The total densities fluctuated seasonally from 200 to 3500 m⁻²; corresponding biomass values ranged from 7 to 600 mg fresh weight m⁻². Maximum density and biomass generally occurred in summer and were minimum in winter. The consistently dominant guild in terms of abundance were the detritivores, mostly comprising Collembola and oribatid mites, accounting for 36–93% of all arthropods; phytophages, predators and detritivores were major guilds in terms of biomass, showing average proportions of 27%, 23% and 20%, respectively. Biomass ratios of predators to prey were generally high in some canopy communities, suggesting the importance of predation pressure in regulating the population levels of arthropods in forests. Average individual size of predators was approximately proportional to that of prey, irrespective of tree locality. Soil arthropod communities maintained densities approximately 10²–10³ times as large as the corresponding canopy communities throughout the year. The seasonal variations in abundance were much greater in canopy than in soil communities.     

Seasonal changes in abundance and spatial distribution of the soil arthropods in a Japanese cedar (Cryptomeria japonica D.Don) plantation,with special reference to collembola and acarina

Seasonal changes in abundance and distribution pattern of soil micro-arthropods were studied in connection with a few environmental factors in a Japanese cedar (Cryptomeria japonica D. Don) plantation. The soil arthropods were sampled from three different depths at intervals of two months for two years. Of the collected animals (total 51000–155000 m⁻²), Collembola (20000–76000 m⁻²), oribatid mites (19000–55000 m⁻²) and carnivorous mites (6200–21000 m⁻²) were the numerically dominant animal groups. Low seasonal variations in abundance indicated their seasonal stability in population levels. The trends in seasonal fluctuation were similar among these groups and between the two years, showing bimodal pattern with little peaks in early summer and winter. The pattern of seasonal fluctuation in abundance of carnivorous mites (P _d) was significantly synchronized with that in the total abundance of Collembola and oribatid mites (P _τ). Thus, the number-ratios (P _d/P _τ) were fairly constant, ranging from 0.10 to 0.25. Seasonal changes in vertical distribution of the three animal groups showed a similar pattern for both years. The downward migrations were shown to be more affected by low temperatures in winter accompanied by snow coverage rather than by the desiccation of the surface soil in summer. All the three groups demonstrated as a whole slightly aggregated patterns of horizontal distribution throughout the two years. Temporal increases in the patchiness indices were observed from summer to autumn when the moisture content of the surface soil was low.     

Effects of mushroom size on the structure of a mycophagous arthropod community: Comparison between infracommunities with different types of resource utilization

The effects of the size of a patchy resource on the structure of a mycophagous arthropod community were examined by comparing numerical responses to the size of individual mushrooms between visitor and dweller communities. A total of 17 fungal genera in nine families were identified during the 13month survey period. A total of 35 arthropod families from nine orders were recorded, with the Hypogastruridae (Collembola) forming 97% of the number of individuals in the visitor community. The response of the visitor community (i.e. number of families, number of individuals, density per mushroom size and diversity) to mushroom size varied in each survey month according to hypogastrurid density. The response of the community composition to mushroom size also varied monthly with hypogastrurid density. In the dweller community, 12 families in three orders were recorded, and one coleopteran and five dipteran families accounted for 90% of the number of individuals. Although the density of dwellers in a mushroom varied between months, the response of the community properties to mushroom size was consistent throughout the survey period. Community composition varied monthly with mushroom size. These results suggest that the size of individual mushrooms is likely to have a greater effect on the structure of dweller communities than visitor communities. Thus, the characteristics of a patchy resource may exert different impacts on the structure of different arthropod communities utilizing the same resource, but in a different manner.     

Abundance patterns of soil micro-arthropods at aPinus pumila scrub in an alpine range of central Japan

Seasonal changes in abundances of major soil micro-arthropods were assessed at aPinus pumila scrub in an alpine range of central Japan during a period with no snow coverage. The total abundance showed a peak in late August, reaching no less than 140 000 m⁻², which was comparable to that in an evergreen coniferous plantation in the cool-temperate zone. Collembola was the most dominant group of soil micro-arthropods, comprising about 50% of the total, followed by oribatid mites (Acari [O]) occupying 20%. Annual mean air temperature was no more than 2.1 °C and the daily fluctuation in temperature was less in soil layers. The thickness of the A₀ layer reached 9–10 cm and soil organic matter accumulation was estimated to be 45–58 ton dry weight ha⁻¹. The large amount of litterfall and organic matter accumulation in the soil, comparable to those of sub-alpine evergreen coniferous forests, and a lower decomposition rate due to severe environmental conditions, suggest the relative importance of litter processing by soil micro-arthropods such as Collembola and Acari, especially in alpine regions.     

An ectomycorrhizal fungus,Cenococcum geophilum,in a coastal pine forest has a high tolerance for an insecticide used to control pine wilt disease

Cenococcum geophilum Fr., one of several ectomycorrhizal species associated with black pine (Pinus thunbergii Parl.), is dominant in the coastal forests of Japan, even under adverse abiotic environmental conditions. In these forests, many tonnes of Sumipine® (fenitrothion) are applied every year to protect P. thunbergii from pine wilt disease, which is transmitted by a beetle. Here, we examined the effect of this insecticide on the species of fungi found as ectomycorrhizae on naturally regenerated P. thunbergii seedlings collected from coastal forest sites that had or had not been sprayed with fenitrothion. The proportion of C. geophilum ectomycorrhizae on black pine root tips was significantly higher in areas where fenitrothion had been applied than in areas where it had not. We measured the in vitro mycelial growth of C. geophilum as well as other ectomycorrhizal fungi of coastal black pine, Rhizopogon roseolus (Corda) Th. Fr. and Pisolithus arhizus (Scop.) Rauschert, at three levels of fenitrothion (density: 1.32 g/cm³), i.e., 0, 0.1 and 0.2 mL L^?1. The growth of all three species decreased significantly as the fenitrothion dosage increased. However, the reduction of mycelial growth in response to fenitrothion was lower in C. geophilum than in the other two species. These results suggest that C. geophilum has a high tolerance for fenitrothion, which may explain its dominance over other ectomycorrhizal species in coastal forests in Japan where fenitrothion is routinely sprayed.